Libytheinae Boisduval et al. 1832

Snout Butterflies

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Introduction

The Libytheinae include fifteen species, two of which are fossils (Kawahara, accepted). Extant species in the subfamily have an unusual global distribution with several widespread species and some that are restricted to insular environments (Shields, 1985a; Kawahara 2006a).

Most species are found in forests or disturbed habitats, but some are also found in dry deserts (Shields 1985a; Kawahara 2006a,b), and at least two species (Libythea labdaca and Libytheana carinenta) are known to migrate in large numbers (Shields, 1987). The male is typically territorial, and both sexes cryptically rest on branches. The number of annual generations is typically two, but this can be affected by environmental factors such as hostplant availability, rainfall, and temperature. The egg is generally laid on buds of Celtis (Celtidaceae), and in nearly all cases, the larva feeds on leaves of this plant genus.

Libytheines can be easily recognized by their exceptionally long labial palpus, although there are a few other nymphalids that have an equally long or longer labial palpus (Ackery et al., 1999). The elongated labial palpus of libytheines is believed to resemble the petiole of a dead leaf, and provide camouflage when the butterfly is at rest (Kawahara, 2003). The foreleg of the libytheine adult male is reduced, but the forleg of the female is well developed. Due to this characteristic, the Libytheinae have traditionally been treated as a separate family (e.g., Pagenstecher, 1901; 1902; 1911), or placed at the base of the Nymphalidae (e.g., Kristensen, 1976; Ackery, 1984; de Jong et al. 1996; Freitas and Brown, 2004). Recent molecular evidence also suggests a basal position of the Libytheinae with respect to the remaining Nymphalidae (Brower, 2000; Wahlberg et al. 2003; 2005).

Characteristics

At least two characters can be used to distinguish the Libytheinae from other Nymphalidae: foreleg reduced in male, but not female; lobe on ventral surface of patagium (Kawahara, accepted).

Discussion of Phylogenetic Relationships

A preliminary phylogenetic analysis of the Libytheinae was conducted (Kawahara, 2001). The parsimony analysis was based on morphological characters for eleven species of Libytheinae and two outgroups. One most parsimonious cladogram was obtained, supporting the monophyly of the subfamily and each genus in the following patterns of species-level relationships (outgroup ((Libytheana carineneta + L. fulvescens)(L. terena + L. motya))(Libythea collenettei (L. myrrha + L. celtis (L. geoffroyi + L. narina)(L. labdaca + L. laius)))). This analysis was preliminary, and the first author is currently reanalyzing the data with additional taxa and characters.

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